Lasius umbratus

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Lasius umbratus
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Tribe: Lasiini
Genus: Lasius
Section: flavus clade
Species group: umbratus
Species: L. umbratus
Binomial name
Lasius umbratus
(Nylander, 1846)

Lasius umbratus casent0005409 profile 1.jpg

Lasius umbratus casent0005409 dorsal 1.jpg

Specimen labels


Common Name
Language: Japanese

Lasius umbratus nests in moist soil in open woodlands, early successional forests, forest edges, and in farm fields (Ellison et al., 2012). It is known to be a temporary social parasite. Queens found new colonies by infiltrating an established nest of another Lasius species (including Lasius alienus (unconfirmed), Lasius brunneus, Lasius emarginatus (unconfirmed), Lasius neoniger, Lasius niger, Lasius pallitarsis, Lasius platythorax and Lasius psammophilus (unconfirmed)), killing the queen and using host workers to care for her initial brood.

At a Glance • Temporary parasite  

Photo Gallery

  • Worker with brood from Harvard, Massachusetts. Photo by Tom Murray.
  • Dealate queen from Groton, Massachusetts. Photo by Tom Murray.
  • Dealate queen.


Collingwood (1979). Clear yellow to reddish yellow; funiculus segments 2 to 4 slightly longer than wide. Scape elliptical in cross section. Petiole tapering to dorsal crest which is usually emarginate. Body surface and appendages covered in adpressed silvery pubescence. Longest hairs on gaster 0.06 mm to 0.11 mm, about half maximum hind tibial width. Erect hairs on genae, scapes and tibiae numerous. Length: 3.8-5.5 mm

Mackay and Mackay (2002) - The workers of this species are yellow or pale brown ants, which have small eyes (35 - 65 ommatidia). The length of the longest hairs on the posterior half of the first gastral tergum (excluding those along the posterior strip) are less than ½ the maximum width of the posterior tibia at mid-length. The hairs on the first gastral tergum are spaced far enough apart that the distances between the tips of most or all hairs are greater than their lengths.

Ellison et al. (2012) - Lasius umbratus is most easily confused with the similarly hairy Lasius subumbratus. However, the hairs on the gaster of L. umbratus are short and bristly – less than half as long as the hind tibia is wide – whereas the hairs on the gaster of L. subumbratus are longer than the hind tibia is wide. In addition, L. umbratus has a concave-topped petiole (viewed from the front or back), whereas L. subumbratus has a convex-topped petiole. An undescribed species of Lasius from New England is similar to L. umbratus except that the new species has erect hairs on its antennal scape and hind tibiae, whereas L. umbratus does not.

Keys including this Species


Widespread over both Eurasia and North America. In eastern North America Lasius umbratus occurs from Nova Scotia south to the Gulf States. Westward, umbratus is abundant through North Dakota, as evidenced by the large numbers of collections made in many localities in that state by G. C. Wheeler and his students. It appears to be relatively common in the southern Rockies, but sparse to absent over most of the rest of western North America.

Latitudinal Distribution Pattern

Latitudinal Range: 65.256706° to 31.156163°.

Tropical South

Distribution based on Regional Taxon Lists

Neotropical Region: Mexico.
Palaearctic Region: Albania, Armenia, Austria, Belarus, Belgium, Bulgaria, China, Croatia, Czech Republic, Democratic Peoples Republic of Korea, Denmark, Estonia, Finland (type locality), Georgia, Germany, Greece, Hungary, Iberian Peninsula, Japan, Latvia, Lithuania, Luxembourg, Montenegro, Netherlands, Norway, Poland, Portugal, Republic of Korea, North Macedonia, Republic of Moldova, Romania, Russian Federation, Slovakia, Slovenia, Spain, Sweden, Switzerland, Türkiye, United Kingdom of Great Britain and Northern Ireland.

Distribution based on AntMaps


Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.


Wilson (1955) - In the face of the revelation that umbratus has a common and hitherto poorly known Palaearctic sibling, Lasius rabaudi (=meridionalis Bondroit), the great mass of European literature pertaining to this species and its many synonyms cannot be accepted without major qualifications. It is in fact very probable that much of the literature deals with rabaudi instead of umbratus. Among the European authors, only Starcke (1937) seems to have fully realized the status and common occurrence of rabaudi and taken this taxonomic information into account in his ecological work. We are still very much in the dark as to whether the two species differ ecologically to any appreciable extent. In the following brief resume, reference to European literature on umbratus is made with the understanding that both species may be included.

In Europe, according to Gosswald (1932), Zimmermann (1934), Donisthorpe (1927), and others, umbratus is less common than the prominent members of Lasius s. s. It prefers dry areas and rarely nests in moist soil; Skwarra (1929) found it very rare on the wet Zehlau moors of East Prussia. It usually nests under rocks, but also occurs in rotting wood (including the timbers of houses), at the foot of trees, or in open ground. On occasion it builds mounds. Its preferred habitat is woodland, but it has also been taken along forest borders and in cultivated fields. In North America, where no sibling comparable to rabaudi is yet known, umbratus differs from the European population in that it prefers moist soil, but it still shows the same latitude in specific nesting sites. The majority of colonies have been taken under stones, while the rest have been taken in or about rotting logs and stumps. I do not know of any case of this ant building mounds or even nesting in the open soil in North America, as it (or rabaudi) has been known to do in Europe. In the northern U. S. east of the Mississippi River, umbratus is limited mostly to moist woodland, where it occurs under a wide variety of conditions of soil texture and insolation. In Alabama and Florida, at the southern extremity of the range, all of the several colonies recorded were found in rotting logs and stumps in swampland. In the western U.S., all of the collections with ecological data that I have examined were made under rocks in open forest and along or near forest borders. In New Mexico, A. C. Cole took this species between 7100 and 8000 feet, always under rocks but under variable conditions of soil moisture and vegetation, e.g. dry soil with scattered juniper and pine or oak and pine, moist soil in a clearing near a hardwood forest, and moist soil in an open grassy area.

As is the case in other species of Lasius, observations on the food habits of umbratus are entirely fragmentary and anecdotal. umbratus is generally thought to be subterranean and to subsist primarily on the excreta of aphids and coccids, since these insects are often found in great numbers in the galleries with the ants (cf. Donisthorpe, 1927, and Gregg, 1944). However, in Holland, Starcke (1937) has observed workers foraging aboveground at night and carrying insects to the nests presumably for use as food. Brown (pers. commun.) has also observed workers aboveground on cloudy days in Pennsylvania.

A number of nuptial flights recorded by Donisthorpe (1927), Crawley (1915), and Eidmann (1926) suggest a long flight season in Europe, extending from as early as August 8 (Crawley) to as late as October 7 (Eidmann). However, there is again no way of knowing whether these records might not represent the overlapping periods of the two species umbratus and rabaudi. Winged queens, determined by me as authentic umbratus, have been taken in Europe on the following dates: IV-4, V-11, VI-9, VI-12, VII-24, VII-31, VIII, VIII, VIII- (15-20), VIII-22, VIII-28, IX3, IX-10, IX-13, IX-16, IX-17, IX-29; these do not involve any apparent geographic trend and by themselves may indicate an unusually long flight period.

The situation in North America is somewhat similar. I have observed queens in flight in the environs of Boston, Mass., in September during two recent seasons. Lone dealate queens were found wandering above ground at Cambridge, Mass., on September 5, 1952, and October 2, 1953, and at Plantagenet, Ontario, on June 30, 1952. Dates on which winged forms have been taken alone or in nido cover the same period, as shown by the following randqm sample: VI-28, VII-8, VII-27, VIII-I, VIII-8, VIII-13, VIII-13, VIII-18, VIII-20, VIII-31, IX-I, IX-5, IX-9, IX-27, X-14, X-28, X-29. Since there are no known sibling species to complicate the picture in North America, the data here suggest that on this continent at least umbratus has an unusually long nuptial season.

There is no evidence to indicate that the reproductives of umbratus build aerial swarms during their nuptial flights, as do those of Lasius niger and Lasius brevicornis, although this does not preclude the possibility. Eidmann (1926) observed queens of umbratus (or rabaudi?) flying singly in Germany, and I have observed definitely determined umbratus queens flying singly on two occasions in the Boston area.

Donisthorpe (1927), Crawley (in Donisthorpe, ibid.), Gosswald (1938), and Holldobler (1953) have reported in detail on the colony founding behavior of "umbratus" and "Lasius mixtus". The normal hosts are niger and Lasius alienus. Under both field and laboratory conditions dealate umbratus queens attack host workers as they encounter them away from the nests, seizing them up the mandibles and carrying them about as they resume foraging. Their victims are usually killed by this treatment and may eventually be eaten. With the fulfillment of this Mordinstinkt, as Holldobler calls it, and the presumed acquisition of the host odor, the queens are ready and able to enter host colonies, although they may be subjected to further attack before acquiring final acceptance. Unfortunately, the authors who have witnessed this phenomenon failed to make a convincing distinction between umbratus and rabaudi, and specimens were not saved to allow corroborative determinations during the present revision.

During the falls of 1952 and 1953 I collected numerous dealate umbratus queens at Cambridge, Mass., and tried introducing them into colonies and colony fragments of Lasius pallitarsis, alienus, and neoniger, but never obtained a complete adoption and saw no evidence of the Mordinstinkt behavior. I have also worked on the theory that the umbratus may join recently fecundated host queens, since both host and parasite queens are often found in species-pure groups under rocks following nuptial flights. Various attempts to bring umbratus and neoniqer queens together, including placing them in the same chamber while chilled, have so far failed; the reason may be, however, that neoniger is not a natural host. I would like to suggest, on what admittedly constitutes negative evidence, that behavior in the population I studied may differ from that in the European populations. If true, this could be due either to geographic variation or to the fact that the European authors were using rabaudi instead of umbratus. Only additional research accompanied by careful determinations will settle the matter.

niger and alienus probably serve as hosts of umbratus in North America as they do in Europe. I have seen two mixed niger-umbratus nest series from Ute Park, New Mexico (A. C. Cole leg. and Coll., MCZ) and one alienus-umbratus series from Beatty, Pennsylvania (Schmitt leg.; MCZ). The Beatty umbratus are minimas. Buren (1944) found a single dealate “aphidicola" queen with a depauperate colony of "flavus nearcticus”, but to my knowledge there has been no additional evidence forthcoming that this or any other Chthonolasius uses species of Cautolasius as hosts.

Collingwood (1979) reporting on umbratus in Europe - This species nests under boulders, in tree stumps and at the base of old trees. Workers are subterranean and seldom or never seen above ground. Flight period from mid August to late September. Single queens found colonies by invasion of and adoption in Lasius niger, Lasius alienus or occasionally Lasius brunneus nests. In late summer dealate queens often wander over the surface of L. niger nests, sometimes carrying a dead L. niger worker as a prelude to securing adoption.

In New Mexico (Mackay and Mackay 2002) - Moist areas, ranging from pastures to mixed forests and pine forests. This species nests under stones and logs, tends Homoptera, and is a temporary social parasite of Lasius alienus, Lasius niger and Lasius neoniger. Reproductives were found in nests July and September, dealate females were found in September. It may form a plesiobiotic relationship with Temnothorax rugatulus.

Association with Other Organisms

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  • This species is a host for the ant Lasius fuliginosus (a temporary parasite) (de la Mora et al., 2021; Janda et al., 2004; Seifert, 2018).
  • This species is a host for the ant Lasius interjectus (a temporary parasite) (de la Mora et al., 2021; Raczkowski & Luque, 2011).
  • This species is a xenobiont for the ant Formica sanguinea (a xenobiont) in Poland (Czechowski & Rotkiewicz, 1997; Kanizsai et al., 2013) (Clearings in a pine forest. In the sandy soil.).
  • This species is a xenobiont for the ant Lasius sabularum (a xenobiont) in Poland (Borowiec, 2011; Kanizsai et al., 2013) (Stand of oak trees. Under stone.).
  • This species is a xenobiont for the ant Polyergus rufescens (a xenobiont) in Poland (Czechowski & Rotkiewicz, 1997; Kanizsai et al., 2013) (Clearings in a pine forest. In the sandy soil.).
  • This species is a host for the cricket Myrmecophilus pergandei (a myrmecophile) in United States.
  • This species is a host for the mite Imparipes obsoletus (a parasite) (Khaustov, 2015) (ectoparasite).
  • This species is a host for the mite Scutacarus longisetus (a parasite) (Khaustov, 2015) (ectoparasite).
  • This species is a host for the phorid fly Pseudacteon formicarum (a parasitoid) (Quevillon, 2018) (encounter mode primary; direct transmission; transmission outside nest). (Quevillon, 2018)
  • This species is a host for the fungus Aegeritella tuberculata (Espadaler & Santamaria, 2012). (Espadaler & Santamaria, 2012)

Flight Period

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec




MCZ-ENT00670481 Lasius umbratus worker hef.jpgMCZ-ENT00670481 Lasius umbratus worker hal.jpgMCZ-ENT00670481 Lasius umbratus worker had.jpgMCZ-ENT00670481 Lasius umbratus worker lbs.JPG
Worker. . Owned by Museum of Comparative Zoology.

Images from AntWeb

Lasius umbratus casent0103993 head 1.jpgLasius umbratus casent0103993 profile 1.jpgLasius umbratus casent0103993 dorsal 1.jpg
Worker. Specimen code casent0103993. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ABS, Lake Placid, FL, USA.
Lasius umbratus casent0172706 head 1.jpgLasius umbratus casent0172706 profile 1.jpgLasius umbratus casent0172706 dorsal 1.jpgLasius umbratus casent0172706 label 1.jpg
Worker. Specimen code casent0172706. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


MCZ-ENT00671591 Lasius umbratus queen hef.jpgMCZ-ENT00671591 Lasius umbratus queen had.jpgMCZ-ENT00671591 Lasius umbratus queen had.jpgMCZ-ENT00671591 Lasius umbratus queen lbs.JPG
Queen. . Owned by Museum of Comparative Zoology.

Images from AntWeb

Lasius umbratus casent0103997 head 1.jpgLasius umbratus casent0103997 profile 1.jpgLasius umbratus casent0103997 dorsal 1.jpgLasius umbratus casent0103997 label 1.jpg
Queen (alate/dealate). Specimen code casent0103997. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ABS, Lake Placid, FL, USA.
Lasius umbratus casent0173146 head 1.jpgLasius umbratus casent0173146 profile 1.jpgLasius umbratus casent0173146 dorsal 1.jpgLasius umbratus casent0173146 label 1.jpg
Queen (alate/dealate). Specimen code casent0173146. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


Images from AntWeb

Lasius umbratus casent0103996 head 1.jpgLasius umbratus casent0103996 profile 1.jpgLasius umbratus casent0103996 profile 2.jpgLasius umbratus casent0103996 dorsal 1.jpgLasius umbratus casent0103996 label 1.jpg
Male (alate). Specimen code casent0103996. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ABS, Lake Placid, FL, USA.
Lasius umbratus casent0173145 head 1.jpgLasius umbratus casent0173145 profile 1.jpgLasius umbratus casent0173145 profile 2.jpgLasius umbratus casent0173145 profile 3.jpgLasius umbratus casent0173145 dorsal 1.jpgLasius umbratus casent0173145 label 1.jpg
Male (alate). Specimen code casent0173145. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by CAS, San Francisco, CA, USA.


The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • umbratus. Formica umbrata Nylander, 1846b: 1048 (q.m.) FINLAND. Schenck, 1852: 59 (w.); Hauschteck, 1962: 219 (k.); Hung, 1969: 456 (k.). Combination in Lasius: Mayr, 1861: 50; in Donisthorpea: Donisthorpe, 1915d: 223; in Formicina: Emery, 1916b: 241; in Acanthomyops: Ruzsky, 1925b: 44; in Chthonolasius: Ruzsky, 1936: 91; in Lasius (Chthonolasius): Ruzsky, 1914a: 59; Müller, 1923: 129; Emery, 1925b: 233; Wilson, 1955a: 150. Senior synonym of belgarum: Wilson, 1955a: 150; Seifert, 1988a: 150; of hirtiscapus: Wilson, 1955a: 150; Seifert, 1990: 11; of affinoumbratus, aphidicola, epinotalis, exacutus, nyaradi, osakana, silvestrii and material of the unavailable names mixtoaffinis, mixtobicornis referred here: Wilson, 1955a: 150; of mixtoumbratus: Wilson, 1955a: 150; Kutter, 1977c: 14. See also: Stitz, 1939: 291; Bernard, 1967: 363; Kutter, 1977c: 232; Yamauchi, 1979: 168; Seifert, 1988a: 150; Atanassov & Dlussky, 1992: 245; Radchenko, 2007: 34.
  • exacutus. Lasius umbratus var. exacutus Ruzsky, 1902d: 15 (w.) RUSSIA. Combination in Acanthomyops: Ruzsky, 1925b: 44; in Lasius (Chthonolasius): Ruzsky, 1914a: 61; Emery, 1925b: 234. Subspecies of umbratus: Ruzsky, 1905b: 287. Raised to species: Ruzsky, 1925b: 44. Subspecies of bicornis: Menozzi, 1939a: 314 (in key). Junior synonym of umbratus: Wilson, 1955a: 150.
  • affinoumbratus. Lasius umbratus var. affinoumbratus Donisthorpe, 1914: 40 (w.) GREAT BRITAIN. Junior synonym of umbratus: Wilson, 1955a: 151.
  • belgarum. Formicina belgarum Bondroit, 1918: 31, fig. 17 (w.q.) BELGIUM. Combination in Lasius (Chthonolasius): Emery, 1925b: 234. Subspecies of umbratus: Stärcke, 1937: 57; of mixtus: Stitz, 1939: 299. Junior synonym of umbratus: Wilson, 1955a: 151; Seifert, 1988: 150.
  • silvestrii. Lasius (Chthonolasius) silvestrii Wheeler, W.M. 1928d: 120 (q.) JAPAN. Junior synonym of umbratus: Wilson, 1955a: 152.
  • hirtiscapus. Lasius umbratus var. hirtiscapus Stärcke, 1937: 43 (q.) CZECHOSLOVAKIA. Subspecies of mixtus: Stitz, 1939: 299. Junior synonym of umbratus: Wilson, 1955a: 152.
  • osakana. Lasius (Chthonolasius) silvestrii var. osakana Santschi, 1941: 278 (q.) JAPAN. Junior synonym of umbratus: Wilson, 1955a: 152.
  • nyaradi. Chthonolasius affinis var. nyaradi Röszler, 1943: 47 (w.q.) ROMANIA. Junior synonym of umbratus: Wilson, 1955a: 152.

Type Material

Wilson (1955) - LECTOTYPE. A dealate queen in the Helsinki Museum, selection by Starcke (1937). Unless otherwise noted the text for the remainder of this section is reported from the publication that includes the original description.



Wilson (1955) - (1) Pilosity and pubescence as in queen. Maximum length of hairs of first gastric tergite anterior to the extreme posterior strip 0.06-0.08 mm., not exceeding one-half the maximum width of the hind tibia at midlength. Alitruncal and cephalic hairs with maximum length of about 0.11 mm.

(2) Petiole in frontal view tapering slightly from the widest point, just above the foramina, to the dorsal crest. The dorsal crest broad and very variable in outline, from flat or even feebly convex to deeply concave; the emargination rounded or angular, never as deep as in Lasius bicornis, i.e. the width (taken at the midpoint of the depth measurement) always exceeds the depth. Intranidal variation considerable; the petiole in a single series may range from flat to distinctly emarginate. In side view the scale is relatively narrow, its dorsal crest acute.

Dong (2017) - (Korean individuals) Body length 4~4.5mm, bright yellow ground color, and body largely hairy. Compound eyes relatively well-developed. Antennal scape protruded head occipital margin in full-face view. Antennal scape with appressed simple hairs densely. Anterior clypeal margin convex, not concaved. Thorax dorsally hairy. Propodeal slope straight and steep. Metapleural gland and propodeal spiracle relatively wide but partially obscured by a couple of setae. In profile view, petiole reversed-V shaped and midpoint of petiole slightly concave or straight when seen in frontal view. Dorsum of petiole with 7~10 filiform fine hairs.


Wilson (1955) - (1) Most of the body surface covered with abundant, relatively short, silvery yellow, predominantly erect hairs. The hairs on the first three gastric tergites with maximum length variable internidally, 0.05-0.11 mm., never more than one-half the maximum width of the hind tibiae at midlength, and often less than one-third; very variable in density, never less than 20 hairs visible above the dorsal profile of the first gastric segment seen in perfect side view and usually more than 30, but never dense enough to overlap one another extensively, Erect hairs forming a fringe on the dorsal crest of the petiole, their maximum length close to that of the gastric hairs. The longest hairs of the alitrunk are on the scutellum, maximum length 0.12-0.21 mm. Maximum length of scutal hairs 0.05-0.15 mm. (see also under geographic variation). Maximum length of cephalic hairs exclusive of those on the clypeus 0.09.-0.11 mm. Body hairs mostly straight or feebly curved, rarely strongly curved (on propodeum) and never sinuate. Standing hairs mayor may not be present on the appendages (see also under geographic variation). All of body and appendages densely covered with short, whitish pubescence which is completely appressed on the body and appressed to decumbent on the appendages; on the gaster it is often abundant enough to obscure partly the shining cuticular surface and to present a whitish overcast to the naked eye.

(2) HW ranging 1.40-1.82 mm. in 45 nest series measured (see also under geographic variation). SI of size extremes 75 and 81.

(3) Petiole in frontal view tapering gradually but distinctly from the level of maximum width (just above the foramina) to the dorsal crest, the width just ventral to the dorsolateral angles 0.9 X the maximum width or less but the frontal outline rarely subquadrate as in rabaudi. Dorsal crest extremely variable in shape, from very feebly concave or even straight to deeply concave with the emargination almost right-angular. The dorsolateral angles always broadly rounded. In side view the scale is narrow and with an acute dorsal crest.

(4) The scape short-elliptical to circular in cross-section, never conspicuously flattened, the minimum width at the midpoint 0.11 mm. or greater. The third funicular segment 1.0-1.5 X longer than broad.

(5) Body medium to dark brown, the appendages lighter, light to medium brown.


Wilson (1955) - (1) Pilosity and pubescence essentially the same as in the queen and worker, except that hairs of the first three gastric tergites are more frequently subdecumbent-suberect. Despite this greater tendency toward obliqueness, the hairs of the first gastric tergite are still too sparse to show much overlap, and their maximum length (excluding those on the extreme posterior strips) ranges internidally 0.07-0.08 mm., or always less than 0.7 X the maximum width of the hind tibia at its midlength. Maximum length of scutellar and cephalic hairs (excluding those on the clypeus) 0.09 mm.

(2) HW 0.85-1.23 mm. in 15 nest series measured; SI of the extremes 62 and 66.

(3) Petiole in frontal view tapering dorsally as in queen and worker. Dorsal margin flat to deeply emarginate, the emargination rounded or angular, never greater than semicircular or rightangular. The scale in side view relatively thin, with an acute dorsal crest.


  • n = 15, 2n = 30 (Switzerland; USA) (Hauschteck, 1962; Hung, 1969).


References based on Global Ant Biodiversity Informatics

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  • Aldawood AS, Sharaf MR (2011) Monomorium dryhimi sp. n., a new ant species (Hymenoptera, Formicidae) of the M. monomorium group from Saudi Arabia, with a key to the Arabian Monomorium monomorium-group. ZooKeys 106: 47–54. doi: 10.3897/zookeys.106.139
  • AntArea. Accessed on February 5th 2014 at
  • Antarea (Personal Communication - Rumsais Blatrix- 27 April 2018)
  • Antarea (at on June 11th 2017)
  • ArtDatabanken Bugs (via GBIG)
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  • Barrett K. E. J. 1970. Ants in France, 1968-69. Entomologist 103: 270-274.
  • Baugnee J. Y. 2003. Camponotus piceus (Leach, 1825), fourmi nouvelle pour la faune belge decouverte dans le parc naturel Viroin-Hermeton (Hymenoptera: Formicidae). Bulletin S. R. B. E./K. B. V. E. 139: 219-225.
  • Behr D., S. Lippke, and K. Colln. 1996. Zur kenntnis der ameisen von Koln (Hymenoptera, Formicidae). Decheniana-Beihefte (Bonn) 35: 215-232.
  • Behr D., and K. Colln. 1993. Zur ameisenfauna (Hymenoptera, Formicidae) von Gonnersdorf (Kr. Daun). Dendrocopos 20: 148-160.
  • Bernard F. 1959. Les fourmis de l'île de Port-Cros. Contribution à l'écologie des anciennes forêts méditerranéennes. Vie Milieu 9: 340-360.
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  • Bernard F. 1976. Écologie des fourmis des grès d'Annot, comparées à celles de la Provence calcaire. Annales du Muséum d'Histoire Naturelle de Nice 3: 33-54.
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